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南亚热带森林生态系统磷循环及其对全球变化的响应

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  ByWenjuanHuangADissertationSubmittedtoGraduateUniversityofChineseAcademyofSciencesInpartialfulfillmentoftherequirementForthedegreeofDoctorofEcologySouthChinaBotanicalGardenChineseAcademyofSciencesMay2013

  致谢致谢lNilIII[IIIIllIllllllllMlUlIIIIIllIY2368528六年前,怀着对科研的崇敬与热爱走进华南植物园的大门,开始了一段新的人生旅程。时光如梭,悄然而逝。回首走过的六年科研生活,我懵懂过、无知过、努力过,渐渐地成熟。首先,我要感谢我的导师周国逸研究员。刚到华南植物园,周老师的平易近人、顾盼神飞、风趣幽默给我留下了深刻的印象。一路走来,曾记得周老师和我讲过许多人生道理,在科研之路上不断地指引着我前进的方向:曾记得周老师给我讲述生态学的知识,让我感受到了一个大科学家的思想;曾记得周老师对我慈父般的教诲,让我深刻体会到“天才是百分之九十九的勤奋加百分之一的灵感”;曾记得每当我的文章被接收时周老师同我一起激动与开心,让我懂得了老师与学生的心连心。这些年,“师恩如父,恩重如山”已深深地种在了我的心里。我的另一位导师刘菊秀副研究员,是我的良师益友。科研之路,是刘老师牵着我的手陪我一起走过。我的每一次报告,刘老师都认真细心地指导:我的每一篇论文,刘老师都逐字逐句地修改。刘老师的那份睿智、踏实、严谨、认真每时每刻都感染着我。在我迷茫的时候,是刘老师让我再次坚定了信念;在我失落的时候,是刘老师给我力量让我勇敢地走下去;在我收获的时候,是刘老师为我喝彩。我俩师生之情,就像一杯酒,一首老歌。能遇到刘老师这样的好老师是我的荣幸。在六年的学习生活中,我也得到了课题组各位老师的指导和帮助。张德强老师和蔼的笑容和一颗总为学生着想的心,总让我不时地感觉到课题组的温暖。还记得李跃林老师彻夜地在帮我修改我的第一篇英文文章;还记得唐旭利老师在我刚入园时给予我大量的帮助与指导;还记得闫俊华老师在我的多次报告中提出的宝贵意见。谢谢你们!感谢张倩媚老师和龚文璇女士在学习的日常事务中给予的支持与帮助。感谢刘世忠老师和褚国伟老师在野外采样过程中的鼎力相助。感谢华南植物园实验室的余清发老师、李红英女士和赖卓银女士在实验室分析过程中给予的指导与帮助。在GriffithUniversity的学习中,感谢ZhihongXu老师在学习与生活对我的指导与照顾,他的平易近人让我身在异乡也能感受到温暖;感谢ChengrongChen

  南亚热带森林生态系统磷循环及其对全球变化的响应老师对我的实验设计与实验方法的耐心指导,他严谨求实、一丝不苟的精神是我学习的榜样;感谢王应平老师对我英文文章的修改与投稿过程中投入大量的心血,他渊博的知识、敏锐的思维和可爱的笑容给我留下了极其深刻的印象。感谢美国UCDavis的BenjaminZ.Houlton老师在学习上的指导与帮助,他开阔的思维让我深受启发。感谢黄钰辉师兄在我刚入园时对我的照顾,让我很快地适应了广州的生活。感谢刘海桂师姐的关照,与你一起逛街吃饭的日子很开心。感谢段洪浪师兄从大学到研究生一路的帮助,让我少走了许多弯路。感谢李银、韩天丰、李义勇和徐伟强无论在实验还是生活中给予大量无私的帮助,你们的可爱与风趣让我的生活增添了不少色彩。感谢杨方方师姐、江浩师兄、刘兴诏师兄、孙芳芳师姐、王国勤师姐、吴建平师兄、刘申师兄、汤新艺师兄、田晓雪师姐、罗艳师姐等对我学习的帮助与生活的关心。感谢陈小梅、侯恩庆、周艺艽、乔玉娜、张娜、肖盏、梁国华、李坤、李荣华、赵亮、陈青青、黄德卫、刘效东、丘清燕、张灏、张亚茹、郑克举、方熊、苗娟、欧阳旭、张静、龙凤玲等,和你们在一起学习、讨论、走路、吃饭、聊天、打球、打牌、唱歌的日子很美好!感谢GriffithUniversity的朋友刘先、喻乐、芮亦超、段吉闯、徐岩、FrederiqueReverchon、KadumM.Abdullah、滕应教授、马琪女士,与你们在一起一年的日子很快乐!感谢UCDavis的好友AlisonMarklein、SaraEnders、ErinLennon、JoyCookingham、MeaganMnich、ScottMorford、JohnBaum、HannaMorris、JamesFarlin,和你们在一起半年的时光让我感到很开心。感谢赖志敏部长、宋丽英老师和庞文峰老师、余艳老师、陈少薇老师、雷广霖老师和吴卓颜老师在过去的学习、实验以及生活中给予的关心与帮助。感谢邓汝芳、朱飞飞和林国俊,从大学四年的同窗到六年的博士生生活,有你们相伴,让我不再孤单。感谢我的舍友周俊芳、黄柳菁和郝香芬,单调的生活中因你们而多了许多阳光与欢笑。在此,也要感谢我的好友古曦的一路相伴!感谢我的爱人黄少君对我无私的包容与无限的支持!特别感谢我的家人和所有的亲人们多年来对我学业和生活的无私奉献和默默支持!

  摘要南亚热带森林生态系统磷循环及其对全球变化的响应磷(P)的有效性平衡调控着热带亚热带森林生态系统的结构和功能。全球变化,如大气C02浓度升高,大气N沉降增加和降水格局改变,都将对P的生物地球化学循环产生重大的影响。本文以鼎湖山自然保护区三个典型南亚热带地带性植被——季风常绿阔叶林(简称季风林)及其演替系列——针阔叶混交林(简称混交林)和马尾松林为研究对象,通过定位观测和长期实验数据,研究不同演替阶段森林生态系统P的有效性与森林生产力的关系,探讨生态系统P循环随着森林演替的变化情况。同时,我们通过人工控制实验(包括碳氮交互实验、人工模拟N沉降和降水改变),研究全球变化对南亚热带森林生态系统P循环的影响。碳氮交互实验运用大型开顶箱,模拟大气C02浓度升高(700gm01.mol一)和N沉降(10gNm’2yr‘1)增加的情形。人工模拟N沉降包括对照(OgNm一2yr-1)、低N(5gNm。2yr-1)、中N(10gNm。2yr。1)和高N(15gNm一2yr一1)处理。降水改变包括移除降雨、自然降雨(对照)和加倍降雨处理。通过在鼎湖山自然保护区内定位研究和长期观测得出:自1989年以来,该地区大气N沉降达到了4.6gNm‘2yrl。从1978年至2008年,季风林的现存生物量和地上部分净初级生产力都表现出逐渐下降的趋势。马尾松林、混交林和季风林植被水平的叶片氮磷比(N:P)分别为22.5、24.7和25.8,即随着森林正向演替逐渐增大。季风林受高N沉降的影响和系统对N资源的自我积累,土壤中N有效性比马尾松林和混交林高,因此,需要更多的P来满足植物生长的需要。但在季风林中,更多的P被固存于土壤有机质中,导致土壤有效P的储量比马尾松林和混交林的低。该结果表明,N的过量和P的缺乏可能是季风林衰退的原 因。 进一步分析三个森林内主要树种树冠下的土壤磷酸酶活性、有效P含量和植 物体N:P,可得出:在整个群落水平上,马尾松林、混交林和季风林的土壤磷酸 酶活性(O-20 cm)分别为7.2 gmol p—NP g—h~、13.0 gmolp.NP g—h。和16.1 gmol P—NP g。1 h~;演替早期森林的主要树种树冠下的土壤磷酸酶活性或每单位土壤有

  南亚热带森林生态系统磷循环及其对全球变化的响应 机C的磷酸酶活性都较演替后期森林低。结果表明,随着森林正向演替,森林 生态系统受P限制的程度逐渐加大;在季风林,与其它树种相比,占生物量大多 数的主要树种更受P的限制。 通过人工控制实验,研究全球变化对南亚热带森林生态系统P循环的影响结 果表明:C02浓度升高,使土壤湿度加大,有利于土壤有机质的矿化,提高土壤 无机P含量,促进植物对P的吸收,导致某些树种植物体N:P的下降。在C02 浓度升高的背景下,相比于非豆科植物,N添加(10 gNm。2 yrl)更有利于降低 了豆科植物的N:P。植物体N:P的变化主要与其P含量的变化有关。因此,在未 来C02浓度升高和大气N沉降增加的背景下,南亚热带森林植物可以通过加大 对P循环的影响来缓解植物受P限制的问题。但N添加对南亚热带森林生态系 统P循环的影响会因系统不同的N水平而异。在N水平较低的马尾松林和混交 林,适量的N添加可以通过提高土壤磷酸酶活性来提高植物对P的吸收,但对 于N饱和的季风林而言,过量的N添加降低土壤磷酸酶活性,导致植物叶片N:P 的增加,进而加剧系统受P的限制。最后,土壤水分有效性也与土壤磷酸酶活性 有密切的关系。在旱季,三个森林土壤磷酸酶活性随降雨量的增加表现出逐渐增 大的趋势;在雨季,与对照相比,移除降雨处理分别导致马尾松林、混交林和季 风林的土壤磷酸酶活性显著地下降了25%,31 O/o并n 32%,而加倍降雨处理仅显著 地降低了季风林的土壤磷酸酶活性。因此,在南亚热带森林生态系统,全年的干 旱会降低土壤有机P的矿化潜力,而在雨季,降雨量的增加将不利于季风林土壤 有机P的矿化。 本文研究结果表明,在南亚热带地区,由于长期受高N沉降的影响和随森 林正向演替对N资源的自我积累,生态系统受P限制的压力在成熟森林中最大。 C02浓度升高可以通过加大对P循环的影响来缓解南亚热带森林植物生长受P限 制。N沉降增加和降水改变对南亚热带森林生态系统P循环的影响会因不同演替 阶段森林而异。 关键词:南亚热带森林;磷限制;磷酸酶活性;氮磷比;全球变化

  Abstract Phosphorus cycle in subtropical forests and its responses to global change By Wenj uan Huang Directed by Guoyi Zhou and Juxiu Liu Abstract Phosphorus(P)availability is critical to control ecosystem structure and function in tropical and subtropical forests.Global change,such as elevated atmospheric carbon dioxide(COz)concentrations,increased Ndeposition,and precipitation changes,has greatly altered Pcycle.In this studywe selected three typical subtropical forests at Dinghushan Biosphere Reserve(DBR)in southern China.The three forests were pine forest(PF),mixed pine and broadleaf forest(MF)and monsoon evergreen broadleaf forest(MEBF),which represented the earlymiddle and late successional stages of subtropical forests,respectively.Data were collected from the long—term research plots in the forests to study the relationship between Pavailability and forest productivity.The hypothesis was that Plimitation would increase with forest succession.Three controlled experiments(elevated C02 and Naddition,simulated Ndeposition and precipitation changes)were also used to study the effects of global change on Pcycle in subtropical forests.We used model forest ecosystems in open.top chambers to study the effects of elevated C02(ca.700 pmol mol一1)alone and together with Naddition(1 0gNm‘2 yr’’)on Pcycles.The experiment of simulated Ndeposition included four Naddition treatments,which were control(0 gNm‘2 yr一1),low.N(5 gNm一2 yr一1),medium.N(10 gNm一2 yr-1)and high.N(15 gNm叱yr叫).Three treatments(no precipitation,natural precipitation and doubled ,,一一,一一precipitation)were conducted in the precipitation experiment. Total Ndeposition at Dinghushan Biosphere Reserve was about 4.6 gNm‘2 yr4 1after 1989.The standing biomass and productivity of MEBF have been declining over the last 30 years.The ratios of Nand P(N:P)in the living leaves were 22。5 for PF’ 24.7 for MF and 25.8 for MEBFshowing an increase trend with community

  南亚热带森林生态系统磷循环及其对全球变化的响应 succession.Our finding suggests that the higher Nstock in soil pools through atmospheric deposition and self-accumulation in MEBF than in PF and MF would require more available Pto prevent deficiency that would limit plant growth.However as more organic matter accumulated and thus,more Pwas bound in MEBF,there was much less available Pin soils.These processes resulted in significantly higher N:P in living leaves and Pshortage in the old-growth forest,which may be responsible for the decline. We further investigated soil phosphatase activity underneath tree species in PF MF and MEBFas well as soil available Pand Nand Pstoichiometry of the tree species.The results showed that phosphatase activities in bulk soils at the top 20 cm mineral soils were 7.2 gmol p-NP g一1 h~for PF1 3.0 I-tmol p-NP 91 h一1 for MF and 16.1 gmol p-NP g~h-Ifor MEBF.Soil phosphatase activities or soil phosphatase activities per unit soil organic carbon under tree species were relatively low in the early successional forest.Our results imply that Plimitation increases with forest succession.The dominant tree species with low soil phosphatase activity in MEBF are likely more P-limited than other tree species. Results from the control experiments showed that:(1)elevated C02 increased soil inorganic Pdue to the greater soil moisture,and thus,increased Pconcentration and decreased N:P ratios in some plant tissues of tree species.N addition favored the N2 fixer rather than the non.N2 fixers to decrease N:P ratios in plants under elevated C02.The reductions of N:P ratios in response to elevated C02 and Naddition were mainly associated with the increases in Pconcentrations.Our results imply that elevated C02 and Naddition could facilitate tree species to mitigate Plimitation by more strongly influencing on Pdynamics than Nin the subtropical forests.(2) Howeverthe effects of Naddition on Pdynamics were dependent on Navailability in forest ecosystems.The moderate Naddition(10w—N addition)could facilitate Plimitation to plant growth through enhancing soil phosphatase activities in PF and MF while more Naddition to MEBF depressed soil phosphatase activity and increased N:P in living leaves,which would further aggravate Plimitation.(3)Soil phosphatase activity was also closely related to soil water availability.In the dry season,soil

  Abstract phosphatase activity in the three forests showed arising trend with increasing precipitation treatments。In the wet season,soil phosphatase activity in no precipitation was significantly decreased by 25%for PF3 1%for MF and 32%for MEBF when compared with contr01.Doubled precipitation significantly depressed soil phosphatase activity by 15%than control in MEBF in the wet season.Our results suggest that adecrease in organi cPturnover would occur in the three forests if there was adrought in awhole yearwhile more rainfall in the wet season would also be adverse to organic Pturnover in MEBF Therefore,our results indicate that Plimitation increases with forest succession through atmospheric Ndeposition and Nself-accumulation in subtropics.Elevated C02 could favor plant growth by greatly influencing Pdynamics in subtropical forests. The impacts of increased Ndeposition and precipitation changes on Pdynamic are closely related to forest succession in subtropics. Keywords:Subtropical forests,Phosphorus limitation,Phosphatase activityN:P’ Global change

  目录 目录 jlI:谢……………………………………………………………………………………………………………………………….0 摘要………………………………………………………………………………………………………………………………。i Abstract………………………………………………………………………………………………………………………..I 弓I言……………………………………………………………………………………………………………………………….1 第一章、绪论…………………………………………………………………………………3 第一节、生态系统磷循环的基本过程……………………………………………………………3 一、土壤磷的有效性………………………………………………………………………………4 二、植物对磷的吸收………………………………………………………………………………5 三、微生物对磷的影响……………………………………………………………………………6 第二节、森林生态系统的磷限制…………………………………………………………………7 一、森林生态系统磷限制的地域分布特征………………………………………………………7 二、磷限制的指标…………………………………………………………………………………9 l、氮磷比…………………………………………………………………………………………。9 2、磷酸酶活性……………………………………………………………………………………10 第三节、森林演替对森林生态系统磷的影响………………………………………………….12 一、植被初生演替……………………………………………………………………………….12 二、植被次生演替……………………………………………………………………………….13 第四节、全球变化对森林生态系统磷的影响………………………………………………….15 一、二氧化碳浓度升高对森林生态系统磷的影响…………………………………………….15 二、氮沉降对森林生态系统磷的影响………………………………………………………….17 三、降水改变对森林生态系统磷的影响……………………………………………………….19 第五节、结语……………………………………………………………………………………2l 第二章、研究地概况和研究方法……………………………………………。24 第一节、自然林实验……………………………………………………………………………24 一、鼎湖山自然保护区研究地概况…………………………………………………………….24 二、永久样地建立……………………………………………………………………………….26 三、长期监测实验……………………………………………………………………………….26 1、大气氮沉降……………………………………………………………………………………26 2、生物量调查……………………………………………………………………………………26 3、地上部分净初级生产力的估算………………………………………………………………27 4、样品采集………………………………………………………………………………………27 4.1叶片采集……………………………………………………………………………………一27 4.2凋落物采集…………………………………………………………………………………一27 4.3土壤样品采集………………………………………………………………………………..28 四、不同树种对磷有效性的影响实验………………………………………………………….28

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